Evolutionary biologists, evolutionary epistemologists, and biosemioticians have demonstrated that organisms not merely adapt to an external world, but that they actively construct their environmental, sociocultural, and cognitive niches. Denis Noble demonstrates that such is no different for those organisms that engage in science, and he lays bare several crucial assumptions that define the scientific dogmas and practices of evolutionary biology.

This special issue for the Journal for General Philosophy of Science is devoted to exploring the impact and many ramifications of current research in evolutionary epistemology. Evolutionary epistemology is an inter- and multidisciplinary area of research that can be divided into two ever-inclusive research avenues. One research avenue expands on the EEM program and investigates the epistemology of evolution. The other research avenue builds on the EET program and researches the evolution of epistemology. Since its conception, EE has developed three (...) schools of thought: adaptationist, non-adaptationist, and applied EE. Although diverse in outlook and theoretical background, these research avenues and schools share the same agenda of understanding how knowledge evolves, and how it relates to the world. In this paper, we first explain wherefrom evolutionary epistemological schools of thought developed, and then we highlight current debates in EE by briefly reviewing the papers that form part of this special issue. (shrink)

Group-structured populations, of the kind prominent in discussions of multilevel selection, are contrasted with ‘neighbor-structured’ populations. I argue that it is a necessary condition on multilevel description of a selection process that there should be a nonarbitrary division of the population into equivalence classes (or an approximation to this situation). The discussion is focused via comparisons between two famous problem cases involving group structure (altruism and heterozygote advantage) and two neighbor-structured cases that resemble them. Conclusions are also drawn about the (...) role of correlated interaction in the evolution of altruism. 1 Introduction 2 Two Kinds of Population Structure 3 Objections and Replies 4 Particles on a Line 5 Conclusion Appendix: Neighborhoods and Selection CiteULike Connotea Del.icio.us What's this? (shrink)

Both biologists and philosophers often make use of simple verbal formulations of necessary and sufficient conditions for evolution by natural selection (ENS). Such summaries go back to Darwin's Origin of Species (especially the "Recapitulation"), but recent ones are more compact.1 Perhaps the most commonly cited formulation is due to Lewontin.2 These summaries tend to have three or four conditions, where the core requirement is a combination of variation, heredity, and fitness differences. The summaries are employed in several ways. First, they (...) are often used in pedagogical contexts, and in showing the coherence of evolutionary theory in response to attacks from outside biology. Second, they are important in discussions of extensions of evolutionary principles to new domains, such as cultural change. The summaries also have intrinsic scientific and philosophical interest as attempts to capture some core principles of evolutionary theory in a highly concise way. Despite their prominence, both the proper formulation and status of these summaries are unclear. Standard formulations are subject to counterexamples, and their relations to formal models of evolutionary change are not straightforward. Here I look closely at these verbal summaries, and at how they relate to formal models. Are the summaries merely rough approximations that have no theoretical role of their own? Perhaps they could operate as theoretical statements in Darwin's time, but have now been superseded by more exact treatments. (shrink)

Kerr and Godfrey-Smith argue that two mathematically equivalent, alternative formal representations drawn from population genetics, the contextualist and collectivist formalisms, may be equally good for quantifying the dynamics of some natural systems, despite important differences between the formalisms. I draw on constraints on causal representation from Woodward (Making things happen, Oxford University Press, New York, 2003) and Eberhardt and Scheines (Philos Sci 74(5):981–995, 2006) to argue that one or the other formalism will be superior for arbitrary natural systems in which (...) individuals form different types of groups. (shrink)

Among philosophers, controversy over the notion of drift in population genetics is ongoing. This is at least partly because the notion of drift has an ambiguous usage among population geneticists. My goal in this paper is to explicate the causal dimension of drift, to say what causal influences are responsible for the stochasticity in population genetics models. It is commonplace for population genetics to oppose the influence of selection to that of drift, and to consider how the dynamics of populations (...) are altered when each has greater or lesser influence. I define the causes that are referred to as drift when researchers speak this way. Introduction Populations and Variant Types The Cause–Effect Ambiguity of Drift Non-directional Factors in Population Genetics How N ev Is Used in Population Genetics Causal Conceptions of Drift 6.1 The Millstein/Beatty conception of drift 6.2 Rosenberg and Bouchard: Drift as initial conditions NINPICs 7.1 Why drift is instituted by NINPICs 7.2 How NINPICS work 7.3 NINPICs and random sampling 7.4 Independent sampling and effective population size 7.5 Variance in progeny number 7.6 Population effects of NINPICs NINPICs and the Stochastic Character of Selection Theory Conclusion Appendix CiteULike Connotea Del.icio.us What's this? (shrink)

There is a certain metaphor that has enjoyed tremendous longevity in the evolution of ageing literature. According to this metaphor, nature has a certain goal or purpose, the perpetuation of the species, or, alternatively, the reproductive success of the individual. In relation to this goal, the individual organism has a function, job, or task, namely, to breed and, in some species, to raise its brood to maturity. On this picture, those who cannot, or can no longer, reproduce are somehow invisible (...) to, or even dispensable to, the evolutionary process. Here, I argue that the metaphor should be discarded, not on the grounds that it is a metaphor, but on the grounds that this particular metaphor distorts our understanding of the evolution of ageing. One reason the metaphor is problematic is that it frames senescence and death as nature’s verdict on the value of older individuals. Instead, we should explore a different metaphor: the lengthy post-reproductive period in humans and some other animals is not an accident of culture, but designed by nature for the purpose of supporting and guiding younger generations. On this alternate picture, different stages of life have their own evolutionary rationales, their distinctive design features, their special mandates. (shrink)

Over the last several decades Ellen Dissanayake has developed an evolutionary theory of art that views all art as having evolved for the function of promoting group cohesion. This theory is not without its critics, yet it has received little empirical attention. In this article I propose a more modest formulation of Dissanayake’s hypothesis and proceed to test it using a cross-cultural analysis. I rely on the ethnographic databases of the electronic Human Relations Area Files as well as the Standard (...) Cross-Cultural Sample to compare proxies for group cohesion with measures of ritual art, a subset of art that I theoretically define. While my results cannot decisively prove or disprove Dissanayake’s hypothesis, they do lend empirical support to her views. This is promising not only for her views but for outlining ways of empirically testing other views in evolutionary aesthetics. (shrink)

Recent studies in the evolution of cooperation have shifted focus from altruistic to mutualistic cooperation. This change in focus is purported to reveal new explanations for the evolution of prosocial behavior. We argue that the common classification scheme for social behavior used to distinguish between altruistic and mutualistic cooperation is flawed because it fails to take into account dynamically relevant game-theoretic features. This leads some arguments about the evolution of cooperation to conflate dynamical scenarios that differ regarding the basic conditions (...) on the emergence and maintenance of cooperation. We use the tools of evolutionary game theory to increase the resolution of the classification scheme and analyze what evolutionary inferences classifying social behavior can license. (shrink)

Framing systematics as a field consistent with scientific inquiry entails that inferences of phylogenetic hypotheses have the goal of producing accounts of past causal events that explain differentially shared characters among organisms. Linking observations of characters to inferences occurs by way of why-questions implied by data matrices. Because of their form, why-questions require the use of common-cause theories. Such theories in phylogenetic inferences include natural selection and genetic drift. Selection or drift can explain ‘morphological’ characters but selection cannot be causally (...) applied to sequences since fitness differences cannot be directly associated with individual nucleotides or amino acids. The relation of selection to sequence data is by way of downward or top-down causation from those phenotypes upon which selection occurs. The application of phylogenetic inference to explain sequence data is thus restricted to instances where drift is the relevant theory; those nucleotides or amino acids that can be explained via downward causation are precluded from inclusion in the data matrix. The restrictions on the inclusion of sequence data in phylogenetic inferences equally apply to species hypotheses, precluding the more restrictive approach known as DNA barcoding. Not being able to discern drift and selection as relevant causal mechanisms can severely constrain the inclusion and explanations of sequence data. Implications of such exclusion are discussed in relation to the requirement of total evidence. (shrink)

The ‘managed-metabolism’ hypothesis suggests that a ‘cooperation barrier’ must be overcome if self-producing chemical organizations are to undergo the transition from non-life to life. This dynamical barrier prevents un-managed autocatalytic networks of molecular species from individuating into complex, cooperative organizations. The barrier arises because molecular species that could otherwise make significant cooperative contributions to the success of an organization will often not be supported within the organization, and because side reactions and other ‘free-riding’ processes will undermine cooperation. As a result, (...) the barrier seriously impedes the emergence of individuality, complex functionality and the transition to life. This barrier is analogous to the cooperation barrier that also impedes the emergence of complex cooperation at all levels of living organization. As has been shown at other levels of organization, the barrier can be overcome comprehensively by appropriate ‘management’. Management implements a system of evolvable constraints that can overcome the cooperation barrier by ensuring that beneficial co-operators are supported within the organization and by suppressing free-riders. In this way, management can control and manipulate the chemical processes of a collectively autocatalytic organization, producing novel processes that serve the interests of the organization as a whole and that could not arise and persist in an un-managed chemical organization. Management self-organizes because it is able to capture some of the benefits that are produced when its interventions promote cooperation, thereby enhancing productivity. Selection will therefore favour the emergence of managers that take over and manage chemical organizations so as to overcome the cooperation barrier. The managed-metabolism hypothesis demonstrates that if management is to overcome the cooperation barrier comprehensively, its interventions must be digitally coded. In this way, the hypothesis accounts for the two-tiered structure of all living cells in which a digitally-coded genetic apparatus manages an analogically-informed metabolism.GraphicalOpen image in new window. (shrink)

Bas van Fraassen’s Darwinian explanation for the success of science has sparked four decades of discussion, with scientific realists and antirealists alike using biologically inspired reasoning to support their points of view. Based on critical engagement with van Fraassen’s proposal itself and later contributions by Stathis Psillos and K. Brad Wray, we claim that central arguments on both sides of this controversy suffer from an insufficient understanding of Darwinism and its underlying biological concepts. Adding the necessary biological background turns out (...) to subvert the argumentative force of viewing the success of scientific theories as analogous to the behaviour of biotic entities. In conclusion, we sketch more productive ways of viewing the relationship between biology and scientific realism. (shrink)

Multi-level selection can be understood via the Price equation or contextual analysis, which offer incompatible statistical decompositions of evolutionary change into components of group and individual selection. Okasha argued that each approach suffers from problem cases. I introduce further problem cases for the Price approach, arguing that it is appropriate for MLS 2 group selection but not MLS 1. I also show that the problem cases Okasha raises for contextual analysis can be resolved. For some such cases, however, it emerges (...) that there is no determinate answer to the question of how much of the total selective effect was due to group selection compared to individual selection. This suggests that when there is interaction between the effect of group character and individual character, one cannot separate selection into distinct ‘levels’ at all. (shrink)

In multicellular organisms, cells are frequently programmed to die. This makes good sense: cells that fail to, or are no longer playing important roles are eliminated. From the cell’s perspective, this also makes sense, since somatic cells in multicellular organisms require the cooperation of clonal relatives. In unicellular organisms, however, programmed cell death poses a difficult and unresolved evolutionary problem. The empirical evidence for PCD in diverse microbial taxa has spurred debates about what precisely PCD means in the case of (...) unicellular organisms. In this article, we survey the concepts of PCD in the literature and the selective pressures associated with its evolution. We show that definitions of PCD have been almost entirely mechanistic and fail to separate questions concerning what PCD fundamentally is from questions about the kinds of mechanisms that realize PCD. We conclude that an evolutionary definition is best able to distinguish PCD from closely related phenomena. Specifically, we define “true” PCD as an adaptation for death triggered by abiotic or biotic environmental stresses. True PCD is thus not only an evolutionary product but must also have been a target of selection. Apparent PCD resulting from pleiotropy, genetic drift, or trade-offs is not true PCD. We call this “ersatz PCD.”. (shrink)

Clade selection is unpopular with philosophers who otherwise accept multilevel selection theory. Clades cannot reproduce, and reproduction is widely thought necessary for evolution by natural selection, especially of complex adaptations. Using microbial evolutionary processes as heuristics, I argue contrariwise, that (1) clade growth (proliferation of contained species) substitutes for clade reproduction in the evolution of complex adaptation, (2) clade-level properties favoring persistence – species richness, dispersal, divergence, and possibly intraclade cooperation – are not collapsible into species-level traits, (3) such properties (...) can be maintained by selection on clades, and (4) clade selection extends the explanatory power of the theory of evolution. (shrink)

Here I advance two related evolutionary propositions. (1) Natural selection is most often considered to require competition between reproducing “individuals”, sometimes quite broadly conceived, as in cases of clonal, species or multispecies-community selection. But differential survival of non-competing and non-reproducing individuals will also result in increasing frequencies of survival-promoting “adaptations” among survivors, and thus is also a kind of natural selection. (2) Darwinists have challenged the view that the Earth’s biosphere is an evolved global homeostatic system. Since there is only (...) one biosphere, reproductive competition cannot have been involved in selection for such survival-promoting adaptations, they claim. But natural selection through survival could reconcile Gaia with evolutionary theory. (shrink)

Reydon and Scholz raise doubts about the Darwinian status of organizational ecology by arguing that Darwinian principles are not applicable to organizational populations. Although their critique of organizational ecology’s typological essentialism is correct, they go on to reject the Darwinian status of organizational populations. This paper claims that the replicator-interactor distinction raised in modern philosophy of biology but overlooked for discussion by Reydon and Scholz provides a way forward. It is possible to conceptualize evolving Darwinian populations providing that the inheritance (...) mechanism is appropriately specified. By this approach, adaptation and selection are no longer dichotomized, and the evolutionary significance of knowledge transmission is highlighted. (shrink)

Interdisciplinarity is one of the features of modern science, defined as blurring the boundaries of disciplines and overcoming their limitations or excessive specialization by borrowing methods from one discipline into another, integrating different theoretical assumptions, and using the same concepts and terms. Often, theoretical knowledge of one discipline and technological advances of another are combined within an interdisciplinary science, and new branches or disciplines may also emerge. Biosemiotics, a field that arose at the crossroads of biology, semiotics, linguistics, and philosophy, (...) enables scientists to borrow theoretical assumptions from semiotics and extend them to different biological theories. The latter applies especially to extended synthesis, wherein culture is viewed as one of the factors influencing evolution. In the present research, the semiotic system of Ukrainian folk ornament is analyzed through the theory of fractals, key features of which are recursion and self-similarity. As a result, an assumption is made about the fractal structure of culture and social life on a conceptual level. What follows is a discussion of how this assumption can contribute to the multilevel selection theory, one of the foundations of extended synthesis, which employs the concept of self-similarity at all levels of the biological hierarchy. (shrink)

Individuality is an important concept in biology, yet there are many non-equivalent criteria of individuality expressed in different kinds of biological individuals. This article evaluates these different kinds in terms of their capacity to support explanatory generalizations over the systems they individuate. Viewing the problem of individuality from this perspective promotes a splitting strategy in which different kinds make different epistemic trade-offs that suit them for different explanatory roles. I argue that evolutionary individuals, interpreted as forming a functional kind, face (...) difficulties of individuation and explanatory power that are mitigated by relying on more structurally based properties and non-evolutionary kinds. 1Introduction2Kinds of Biological Individuals3Evolutionary Individuals and Functional Kinds4Evolutionary versus Non-evolutionary Kinds of Individuals 4.1Physiological individuality4.2Ecological individuality4.3Developmental individuality5Conclusion. (shrink)

Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with (genotypically represented) fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences (...) among genes within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology (evo-devo) during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models (both classical evolutionary theory and evo-devo) on the other. (shrink)

Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences among genes (...) within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models on the other. (shrink)

Recent evolutionary perspectives on guilt tend to focus on how guilt functions as a means for the individual to self-regulate behavior and as a mechanism for reinforcing cooperative tendencies. While these accounts highlight important dimensions of guilt and provide important insights into its evolutionary emergence, they pay scant attention to the large empirical literature on its maladaptive effects on individuals. This paper considers the nature of guilt, explores its biological function, and provides an evolutionary perspective on whether it is an (...) individual-level or group selected trait. After surveying philosophical and psychological analyses of guilt, we consider which psychological mechanisms underlie the capacity to experience and act from guilt and whether they point to an emergence of guilt in early humans or to guilt having a longer phylogenetic history. Because guilt is a characteristically social emotion, we then examine its contemporary role in social and legal contexts, which may p... (shrink)

Several prominent debates in biology, such as those surrounding adaptationism, group selection, and punctuated equilibrium, have focused on disagreements about the relative importance of a cause in producing a phenomenon of interest. Some philosophers, such as John Beatty have expressed scepticism about the scientific value of engaging in these controversies, and Karen Kovaka has suggested that their value might be limited. In this paper, I challenge that scepticism by giving a novel analysis of relative significance controversies, showing that there are (...) three forms they can take. I argue that these controversies can have significant epistemic upshots, in that they help scientists form predictions about new instances of the phenomenon of interest. Finally, using two historical examples, I show how engaging in these controversies can improve our understanding of causal relationships. (shrink)

In the scientific literature on religious evolution, two competing theories appeal to group selection to explain the relationship between religious belief and altruism, or costly, prosocial behavior. Both theories agree that group selection plays an important role in cultural evolution, affecting psychological traits that individuals acquire through social learning. They disagree, however, about whether group selection has also played a role in genetic evolution, affecting traits that are inherited genetically. Recently, Jonathan Haidt has defended the most fully developed account based (...) on genetic group selection, and I argue here that problems with this account reveal good reasons to doubt that genetic group selection has played any important role in human evolution at all. Thus, considering the role of group selection in religious evolution is important not just because of what it reveals about religious psychology and religious evolution, but also because of what it reveals about the role of group selection in human evolution more generally. (shrink)

Evolutionary Transitions in Individuality have been responsible for the major transitions in levels of selection and individuality in natural history, such as the origins of prokaryotic and eukaryotic cells, multicellular organisms, and eusocial insects. The integrated hierarchical organization of life thereby emerged as groups of individuals repeatedly evolved into new and more complex kinds of individuals. The Social Protocell Hypothesis proposes that the integrated hierarchical organization of human culture can also be understood as the outcome of an ETI—one that produced (...) a “cultural organism” from a substrate of socially learned traditions that were contained in growing and dividing social communities. The SPH predicts that a threshold degree of evolutionary individuality would have been achieved by 2.0–2.5 Mya, followed by an increasing degree of evolutionary individuality as the ETI unfolded. We here assess the SPH by applying a battery of criteria—developed to assess evolutionary individuality in biological units—to cultural units across the evolutionary history of Homo. We find an increasing agreement with these criteria, which buttresses the claim that an ETI occurred in the cultural realm. (shrink)

The group selection controversy largely focuses on altruism (e.g., Wilson 1983; Lloyd 2001; Shavit 2004; Okasha 2006, 173ff; Borrello 2010; Leigh 2010; Rosas 2010; Hamilton and Dimond in press). Multilevel selection theory is a resolution of this controversy. Whereas kin selection partitions inclusive fitness into direct and indirect components (via influencing the replication of copies of genes in other individuals), multilevel selection considers within-group and between-group components of fitness (Gardner et al. 2011; Lion et al. 2011). Two scenarios of multilevel (...) selection are often distinguished (Damuth and Heisler 1988; Okasha 2006; Pigliucci 2010): (1) group structure only divides individual fitnesses into within- and between-group components (MLS1); and, (2) groups get their own component of fitness and also, in most definitions, a group-level adaptation (MLS2). (shrink)

In The Secret of Our Success, Joseph Henrich claims that human beings are unique—different from all other animals—because we engage in cumulative cultural evolution. It is the technological and social products of cumulative cultural evolution, not the intrinsic rationality or ‘smartness’ of individual humans, that enable us to live in a huge range of different habitats, and to dominate most of the creatures who share those habitats with us. We are sympathetic to this general view, the latest expression of the (...) ‘California school’s’ view of cultural evolution, and impressed by the lively and interesting way that Henrich handles evidence from anthropology, economics, and many fields of biology. However, because we think it is time for cultural evolutionists to get down to details, this essay review raises questions about Henrich’s analysis of both the cognitive processes and the selection processes that contribute to cumulative cultural evolution. In the former case, we argue that cultural evolutionists need to make more extensive use of cognitive science, and to consider the evidence that mechanisms of cultural learning are products as well as processes of cultural evolution. In the latter case, we ask whether the California school is really serious about selection, or whether it is offering a merely ‘kinetic’ view of cultural evolution, and, assuming the former, outline four potential models of cultural selection that it would be helpful to distinguish more clearly. (shrink)

Darwin’s classic ‘Origin of Species’ (Darwin 1859) described forces of selection acting upon individuals, but there remains a great deal of controversy about what exactly the status and definition of a biological individual is. Recently some authors have argued that the individual is dispensable – that an inability to pin it down is not problematic because little rests on it anyway. The aim of this paper is to show that there is a real problem of biological individuality, and an urgent (...) need to arbitrate over the current plethora of solutions to it. (shrink)

The probability that the fitter of two alleles will increase in frequency in a population goes up as the product of N (the effective population size) and s (the selection coefficient) increases. Discovering the distribution of values for this product across different alleles in different populations is a very important biological task. However, biologists often use the product Ns to define a different concept; they say that drift “dominates” selection or that drift is “stronger than” selection when Ns is much (...) smaller than some threshold quantity (e.g., ½) and that the reverse is true when Ns is much larger than that threshold. We argue that the question of whether drift dominates selection for a single allele in a single population makes no sense. Selection and drift are causes of evolution, but there is no fact of the matter as to which cause is stronger in the evolution of any given allele. (shrink)

The problem of plant individuality is something which has vexed botanists throughout the ages, with fashion swinging back and forth from treating plants as communities of individuals (Darwin 1800 ; Braun and Stone 1853 ; Münch 1938 ) to treating them as organisms in their own right, and although the latter view has dominated mainstream thought most recently (Harper 1977 ; Cook 1985 ; Ariew and Lewontin 2004 ), a lively debate conducted mostly in Scandinavian journals proves that the issues (...) are far from being resolved (Tuomi and Vuorisalo 1989b ; Fagerström 1992 ; Pan and Price 2001 ). In this paper I settle the matter once and for all, by showing which elements of each side are correct. (shrink)

Do multi-level selection explanations of the evolution of social traits deepen the understanding provided by single-level explanations? Central to the former is a mathematical theorem, the multi-level Price decomposition. I build a framework through which to understand the explanatory role of such non-empirical decompositions in scientific practice. Applying this general framework to the present case places two tasks on the agenda. The first task is to distinguish the various ways of suppressing within-collective variation in fitness, and moreover to evaluate their (...) biological interest. I distinguish four such ways: increasing retaliatory capacity, homogenising assortment, and collapsing either fitness structure or character distribution to a mean value. The second task is to discover whether the third term of the Price decomposition measures the effect of any of these hypothetical interventions. On this basis I argue that the multi-level Price decomposition has explanatory value primarily when the sharing-out of collective resources is `subtractable'. Thus its value is more circumscribed than its champions Sober and Wilson (1998) suppose. (shrink)

Microbes are generally thought of as unicellular organisms, but we know that many microbes live as parts of biofilms—complex, surface-attached microbial communities numbering millions of cells. Some authors have recently argued in favour of reconceiving biofilms as biological entities in their own right. In particular, some have claimed that multispecies biofilms are evolutionary individuals : 10126–10132 2015). Against this view, I defend the conservative consensus that selection acts primarily upon microbial cells.

Several recent arguments by philosophers of biology have challenged the traditional view that evolutionary factors, such as drift and selection, are genuine causes of evolutionary outcomes. In the case of drift, advocates of the statistical theory argue that drift is merely the sampling error inherent in the other stochastic processes of evolution and thus denotes a mathematical, rather than causal, feature of populations. This debate has largely centered around one particular model of drift, the Wright–Fisher model, and this has contributed (...) to the plausibility of the statisticalists’ arguments. However, an examination of alternative, predictively inequivalent models shows that drift is a genuine cause that can be manipulated to change population outcomes. This case study illustrates the influence of methodological assumptions on ontological judgments, particularly the pernicious effect of focusing on a particular model at the expense of others and confusing its assumptions and idealizations for true claims about the phenomena being modeled. (shrink)

What changes when an evolutionary transition in individuality takes place? Many different answers have been given, in respect of different cases of actual transition, but some have suggested a general answer: that a major transition is a change in the extent to which selection acts at one hierarchical level rather than another. The current paper evaluates some different ways to develop this general answer as a way to characterise the property ‘evolutionary individuality’; and offers a justification of the option taken (...) in Clarke :413–435, 2013)—to define evolutionary individuality in terms of an object’s capacity to undergo selection at its own level. In addition, I suggest a method by which the property can be measured and argue that a problem which is often considered to be fatal to that method—the problem of ‘cross-level by-products’—can be avoided. (shrink)

Altruism is a deep and complex phenomenon that is analysed by scholars of various disciplines, including psychology, philosophy, biology, evolutionary anthropology and experimental economics. Much confusion arises in current literature because the term altruism covers variable concepts and processes across disciplines. Here we investigate the sense given to altruism when used in different fields and argumentative contexts. We argue that four distinct but related concepts need to be distinguished: (a) psychological altruism , the genuine motivation to improve others’ interests and (...) welfare; (b) reproductive altruism , which involves increasing others’ chances of survival and reproduction at the actor’s expense; (c) behavioural altruism , which involves bearing some cost in the interest of others; and (d) preference altruism , which is a preference for others’ interests. We show how this conceptual clarification permits the identification of overstated claims that stem from an imprecise use of terminology. Distinguishing these four types of altruism will help to solve rhetorical conflicts that currently undermine the interdisciplinary debate about human altruism. (shrink)

Cultural selection models aim to explain cultural phenomena as the products of a selective process, often characterising institutions, practices, norms or behaviours as adaptations. I argue that a lack of attention has been paid to the explanatory power of cultural selection frameworks. Arguments for cultural selection frequently depend on demonstrating only that selection models can in principle be applied to culture, rather than explicitly demonstrating the explanatory payoffs that could arise from their application. Understanding when and how cultural selection generates (...) powerful explanations is crucial to evaluating cultural selection, as well as realising its promised epistemic and practical benefits. I argue that the ability for cultural selection to explain ‘design without a designer’ is crucial to successful and powerful cultural selection explanations. I introduce the strategy of comparing cultural selection to goal-directed agent accounts in order to evaluate when cultural selection can provide distinctive explanatory payoffs, drawing on two case studies to illustrate the benefits of this strategy. I argue that a focus on phenomena which cannot be explained through intention or agency-based explanations in particular could provide a fruitful avenue to identifying the cases where cultural selection can be insightfully applied. (shrink)

References in Darwin’s Origin of Species to competition between units of selection at and above the level of individual organisms are enumerated. In many cases these references clearly speak of natural selection and do not support the view that Darwin thought selection only occurred at the level of the individual organism. Darwin did see organismal selection as the main process by which varieties were created but he also espoused what is here termed community and varietal selection. He saw no essential (...) difference between varieties and species and the references show that he also believed that selection could operate at the species level. (shrink)

In this paper, I describe some strategies for teaching an introductory philosophy of science course to Science, Technology, Engineering, and Mathematics (STEM) students, with reference to my own experience teaching a philosophy of science course in the Fall of 2020. The most important strategy that I advocate is what I call the “Second Philosophy” approach, according to which instructors ought to emphasize that the problems that concern philosophers of science are not manufactured and imposed by philosophers from the outside, but (...) rather are ones that arise internally, during the practice of science itself. To justify this approach, I appeal to considerations from both educational research and the epistemology of testimony. In addition, I defend some distinctive learning goals that philosophy of science instructors ought to adopt when teaching STEM students, which include rectifying empirically well-documented shortcomings in students’ conceptions of the “scientific method” and the “nature of science.” Although my primary focus will be on teaching philosophy of science to STEM students, much of what I propose can be applied to non-philosophy majors generally. Ultimately, as I argue, a successful philosophy of science course for non-philosophy majors must be one that advances a student’s science education. The strategies that I describe and defend here are aimed at precisely that end. (shrink)

In Denis Walsh’s Organisms, Agency, and Evolution, he argues that new developments in the science of biology motivate a radical change to our metaphysical picture of life: what he calls ‘Situated Darwinism’. The central claim is that we should take the biological world to be at base about organisms, and organisms in a fundamentally teleological sense. We critically examine Walsh’s arguments and suggest further developments.

This essay analyzes and develops recent views about explanation in biology. Philosophers of biology have parted with the received deductive-nomological model of scientific explanation primarily by attempting to capture actual biological theorizing and practice. This includes an endorsement of different kinds of explanation (e.g., mathematical and causal-mechanistic), a joint study of discovery and explanation, and an abandonment of models of theory reduction in favor of accounts of explanatory reduction. Of particular current interest are philosophical accounts of complex explanations that appeal (...) to different levels of organismal organization and use contributions from different biological disciplines. The essay lays out one model that views explanatory integration across different disciplines as being structured by scientific problems. I emphasize the philosophical need to take the explanatory aims pursued by different groups of scientists into account, as explanatory aims determine whether different explanations are competing or complementary and govern the dynamics of scientific practice, including interdisciplinary research. I distinguish different kinds of pluralism that philosophers have endorsed in the context of explanation in biology, and draw several implications for science education, especially the need to teach science as an interdisciplinary and dynamic practice guided by scientific problems and explanatory aims. (shrink)

Since van Fraassen first put forward the suggestive idea that many philosophical positions should be construed as ‘stances’ rather than factual beliefs, there have been various attempts to spell out precisely what a philosophical stance might be, and on what basis one should be adopted. In this paper I defend a particular account of stances, the view that they are pragmatically justified perspectives or ways of seeing the world, and compare it to some other accounts that have been offered. In (...) Sect. 2 I consider van Fraassen’s argument for construing empiricism as a stance, and look at some responses to it. In Sect. 3 I outline my conception of stances as perspectives or ways of seeing, and explain how stances so understood may be justified. I illustrate this conception by way of a discussion of the model pluralist position with respect to the units of selection debate in biology, and suggest that on the model pluralist view different perspectives on the units of selection, such as the gene’s eye view, are in fact van Fraassian stances. In Sect. 4 I discuss the view put forward by Teller and Chakravartty among others that stances should be understood as epistemic policies, and argue that it is consistent with the conception of stances as perspectives. In the final section I criticise Rowbottom’s attempt to assimilate stances to Kuhnian paradigms. I argue that he has overlooked some important disanalogies between stances and paradigms, so that the comparison obscures more than it reveals. (shrink)

Evolutionary transitions in individuality (ETIs) are events during which individuals at a given level of organization (particles) interact to form higher-level entities (collectives) which are then recognized as new individuals at that level. ETIs are intimately related to levels of selection, which, following Okasha, can be approached from two different perspectives. One, referred to as ‘synchronic’, asks whether selection occurs at the collective level while the partitioning of particles into collectives is taken for granted. The other, referred to as ‘diachronic’, (...) asks about the origins of the partitioning of particles into collectives. After having presented the two perspectives and a classical formalism used to deal with the levels-of-selection question in the literature, namely the multilevel version of the Price equation, I show that because this formalism treats the levels-of-selection question from a synchronic perspective, it is inadequate to explain ETIs. This is because a fundamental aspect of ETIs is the origin of collectives. From there, I develop a framework for levels of selection compatible with the diachronic perspective. This framework relies on the distinction between what I call, on the one hand, a ‘functional aggregative collective trait’, and on the other hand, a ‘functional non-aggregative collective trait’. After having presented this distinction, I implement it in the Price equation, leading to a new statistical partitioning of this equation which, I argue, represents a causal decomposition more relevant for ETIs. Finally, I exploit this partitioning to explain the critical stages of an ETI. In addition to its explanatory power, a measure of the degree of functional non-aggregativity could be used as a proxy for the degree of individuality of a collective. (shrink)

I argue that Grafen’s formal darwinism project could profitably incorporate a gene’s-eye view, as informed by the major transitions framework. In this, instead of the individual being assumed to maximise its inclusive fitness, genes are assumed to maximise their inclusive fitness. Maximisation of fitness at the individual level is not a straightforward concept because the major transitions framework shows that there are several kinds of biological individual. In addition, individuals have a definable fitness, exhibit individual-level adaptations and arise in a (...) major transition, only to the extent that the inclusive-fitness interests of genes within them coincide. Therefore, as others have suggested, the fundamental level at which fitness is maximised is the gene level. Previous reconciliations of the concepts of gene-level fitness and individual-level fitness implicitly recognise this point. Adaptations always maximise the fitness of their causative genes, but may be simple or complex. Simple adaptations may be controlled by single genes and be maladaptive at higher levels, whereas complex adaptations are controlled by multiple genes and rely on those genes having coinciding fitness interests at a higher level, for a given trait. (shrink)